Minimalist revision and description of 403 new species in 11 subfamilies of Costa Rican braconid parasitoid wasps, including host records for 219 species.

Three new genera are described: Michener (Proteropinae), Bioalfa (Rogadinae), and Hermosomastax (Rogadinae). Keys are given for the New World genera of the following braconid subfamilies: Agathidinae, Braconinae, Cheloninae, Homolobinae, Hormiinae, Ichneutinae, Macrocentrinae, Orgilinae, Proteropinae, Rhysipolinae, and Rogadinae. In these subfamilies 416 species are described or redescribed. Most of the species have been reared and all but 13 are new to science. A consensus sequence of the COI barcodes possessed by each species is employed to diagnose the species, and this approach is justified in the introduction. Most descriptions consist of a lateral or dorsal image of the holotype, a diagnostic COI consensus barcode, the Barcode Index Number (BIN) code with a link to the Barcode of Life Database (BOLD), and the holotype specimen information required by the International Code of Zoological Nomenclature. The following species are treated and those lacking authorship are newly described here with authorship attributable to Sharkey except for the new species of Macrocentrinae which are by Sharkey & van Achterberg: AGATHIDINAE: Aerophiluspaulmarshi, Mesocoelusdavidsmithi, Neothlipsisbobkulai, Plesiocoelusvanachterbergi, Pneumagathiserythrogastra (Cameron, 1905), Therophilusbobwhartoni, T.donaldquickei, T.gracewoodae, T.maetoi, T.montywoodi, T.penteadodiasae, Zacremnopsbrianbrowni, Z.coatlicue Sharkey, 1990, Zacremnopscressoni (Cameron, 1887), Z.ekchuah Sharkey, 1990, Z.josefernandezi, Zelomorphasarahmeierottoae. BRACONINAE: Braconalejandromarini, B.alejandromasisi, B.alexamasisae, B.andresmarini, B.andrewwalshi, B.anniapicadoae, B.anniemoriceae, B.barryhammeli, B.bernardoespinozai, B.carlossanabriai, B.chanchini, B.christophervallei, B.erasmocoronadoi, B.eugeniephillipsae, B.federicomatarritai, B.frankjoycei, B.gerardovegai, B.germanvegai, B.isidrochaconi, B.jimlewisi, B.josejaramilloi, B.juanjoseoviedoi, B.juliodiazi, B.luzmariaromeroae, B.manuelzumbadoi, B.marialuisariasae, B.mariamartachavarriae, B.mariorivasi, B.melissaespinozae, B.nelsonzamorai, B.nicklaphami, B.ninamasisae, B.oliverwalshi, B.paulamarinae, B.rafamoralesi, B.robertofernandezi, B.rogerblancoi, B.ronaldzunigai, B.sigifredomarini, B.tihisiaboshartae, B.wilberthbrizuelai, Digonogastramontylloydi, D.montywoodi, D.motohasegawai, D.natwheelwrighti, D.nickgrishini. CHELONINAE: Adeliusadrianguadamuzi, A.gauldi Shimbori & Shaw, 2019, A.janzeni Shimbori & Shaw, 2019, Ascogastergloriasihezarae, A.grettelvegae, A.guillermopereirai, A.gustavoecheverrii, A.katyvandusenae, A.luisdiegogomezi, Chelonusalejandrozaldivari, C.gustavogutierrezi, C.gustavoinduni, C.harryramirezi, C.hartmanguidoi, C.hazelcambroneroae, C.iangauldi, C.isidrochaconi, C.janecheverriae, C.jeffmilleri, C.jennyphillipsae, C.jeremydewaardi, C.jessiehillae, C.jesusugaldei, C.jimlewisi, C.jimmilleri, C.jimwhitfieldi, C.johanvalerioi, C.johnburnsi, C.johnnoyesi, C.jorgebaltodanoi, C.jorgehernandezi, C.josealfredohernandezi, C.josefernandeztrianai, C.josehernandezcortesi, C.josemanuelperezi, C.josephinerodriguezae, C.juanmatai, C.junkoshimurae, C.kateperezae, C.luciariosae, C.luzmariaromeroae, C.manuelpereirai, C.manuelzumbadoi, C.marianopereirai, C.maribellealvarezae, C.markmetzi, C.markshawi, C.martajimenezae, C.mayrabonillae, C.meganmiltonae, C.melaniamunozae, C.michaelstroudi, C.michellevanderbankae, C.mingfangi, C.minorcarmonai, C.monikaspringerae, C.moniquegilbertae, C.motohasegawai, C.nataliaivanovae, C.nelsonzamorai, C.normwoodleyi, C.osvaldoespinozai, C.pamelacastilloae, C.paulgoldsteini, C.paulhansoni, C.paulheberti, C.petronariosae, C.ramyamanjunathae, C.randallgarciai, C.rebeccakittelae, C.robertoespinozai, C.robertofernandezi, C.rocioecheverriae, C.rodrigogamezi, C.ronaldzunigai, C.rosibelelizondoae, C.rostermoragai, C.ruthfrancoae, C.scottmilleri, C.scottshawi, C.sergioriosi, C.sigifredomarini, C.stevearonsoni, C.stevestroudi, C.sujeevanratnasinghami, C.sureshnaiki, C.torbjornekremi, C.yeimycedenoae, Leptodrepanaalexisae, L.erasmocoronadoi, L.felipechavarriai, L.freddyquesadai, L.gilbertfuentesi, L.manuelriosi, Phanerotomaalmasolisae, P.alvaroherrerai, P.anacordobae, P.anamariamongeae, P.andydeansi, P.angelagonzalezae, P.angelsolisi, P.barryhammeli, P.bernardoespinozai, P.calixtomoragai, P.carolinacanoae, P.christerhanssoni, P.christhompsoni, P.davesmithi, P.davidduthiei, P.dirksteinkei, P.donquickei, P.duniagarciae, P.duvalierbricenoi, P.eddysanchezi, P.eldarayae, P.eliethcantillanoae, P.jenopappi, Pseudophanerotomaalanflemingi, Ps.albanjimenezi, Ps.alejandromarini, Ps.alexsmithi, Ps.allisonbrownae, Ps.bobrobbinsi. HOMOLOBINAE: Exasticolusjennyphillipsae, E.randallgarciai, E.robertofernandezi, E.sigifredomarini, E.tomlewinsoni. HORMIINAE: Hormiusanamariamongeae, H.angelsolisi, H.anniapicadoae, H.arthurchapmani, H.barryhammeli, H.carmenretanae, H.carloswalkeri, H.cesarsuarezi, H.danbrooksi, H.eddysanchezi, H.erikframstadi, H.georgedavisi, H.grettelvegae, H.gustavoinduni, H.hartmanguidoi, H.hectoraritai, H.hesiquiobenitezi, H.irenecanasae, H.isidrochaconi, H.jaygallegosi, H.jimbeachi, H.jimlewisi, H.joelcracrafti, H.johanvalerioi, H.johnburleyi, H.joncoddingtoni, H.jorgecarvajali, H.juanmatai, H.manuelzumbadoi, H.mercedesfosterae, H.modonnellyae, H.nelsonzamorai, H.pamelacastilloae, H.raycypessi, H.ritacolwellae, H.robcolwelli, H.rogerblancosegurai, H.ronaldzunigai, H.russchapmani, H.virginiaferrisae, H.warrenbrighami, H.willsflowersi. ICHNEUTINAE: Oligoneuruskriskrishtalkai, O.jorgejimenezi, Paroligoneuruselainehoaglandae, P.julianhumphriesi, P.mikeiviei. MACROCENTRINAE: Austrozelejorgecampabadali, A.jorgesoberoni, Dolichozelegravitarsis (Muesebeck, 1938), D.josefernandeztrianai, D.josephinerodriguezae, Hymenochaoniakalevikulli, H.kateperezae, H.katherinebaillieae, H.katherineellisonae, H.katyvandusenae, H.kazumifukunagae, H.keithlangdoni, H.keithwillmotti, H.kenjinishidai, H.kimberleysheldonae, H.krisnorvigae, H.lilianamadrigalae, H.lizlangleyae, Macrocentrusfredsingeri, M.geoffbarnardi, M.gregburtoni, M.gretchendailyae, M.grettelvegae, M.gustavogutierrezi, M.hannahjamesae, M.harisridhari, M.hillaryrosnerae, M.hiroshikidonoi, M.iangauldi, M.jennyphillipsae, M.jesseausubeli, M.jessemaysharkae, M.jimwhitfieldi, M.johnbrowni, M.johnburnsi, M.jonathanfranzeni, M.jonathanrosenbergi, M.jorgebaltodanoi, M.lucianocapelli. ORGILINAE: Orgilusamyrossmanae, O.carrolyoonae, O.christhompsoni, O.christinemcmahonae, O.dianalipscombae, O.ebbenielsoni, O.elizabethpennisiae, O.evertlindquisti, O.genestoermeri, O.jamesriegeri, O.jeanmillerae, O.jeffmilleri, O.jerrypowelli, O.jimtiedjei, O.johnlundbergi, O.johnpipolyi, O.jorgellorentei, O.larryspearsi, O.marlinricei, O.mellissaespinozae, O.mikesmithi, O.normplatnicki, O.peterrauchi, O.richardprimacki, O.sandraberriosae, O.sarahmirandae, O.scottmilleri, O.scottmorii, Stantoniabillalleni, S.brookejarvisae, S.donwilsoni, S.erikabjorstromae, S.garywolfi, S.henrikekmani, S.luismirandai, S.miriamzunzae, S.quentinwheeleri, S.robinkazmierae, S.ruthtifferae. PROTEROPINAE: Hebichneutestricolor Sharkey & Wharton, 1994, Proteropsiangauldi, P.vickifunkae, Michenercharlesi. RHYSIPOLINAE: Pseudorhysipolisluisfonsecai, P. mailyngonzalezaeRhysipolisjulioquirosi. ROGADINAE: Aleiodesadrianaradulovae, A.adrianforsythi, A.agnespeelleae, A.alaneaglei, A.alanflemingi, A.alanhalevii, A.alejandromasisi, A.alessandracallejae, A.alexsmithi, A.alfonsopescadori, A.alisundermieri, A.almasolisae, A.alvarougaldei, A.alvaroumanai, A.angelsolisi, A.annhowdenae, A.bobandersoni, A.carolinagodoyae, A.charlieobrieni, A.davefurthi, A.donwhiteheadi, A.doylemckeyi, A.frankhovorei, A.henryhowdeni, A.inga Shimbori & Shaw, 2020, A.johnchemsaki, A.johnkingsolveri, A.gonodontovorus Shimbori & Shaw, 2020, A.manuelzumbadoi, A.mayrabonillae, A.michelledsouzae, A.mikeiviei, A.normwoodleyi, A.pammitchellae, A.pauljohnsoni, A.rosewarnerae, A.steveashei, A.terryerwini, A.willsflowersi, Bioalfapedroleoni, B.alvarougaldei, B.rodrigogamezi, Choreborogasandydeansi, C.eladiocastroi, C.felipechavarriai, C.frankjoycei, Clinocentrusandywarreni, Cl.angelsolisi, Cystomastaxalexhausmanni, Cy.angelagonzalezae, Cy.ayaigarashiae, Hermosomastaxclavifemorus Quicke sp. nov., Heterogamusdonstonei, Pseudoyeliconesbernsweeneyi, Stiropiusbencrairi, S.berndkerni, S.edgargutierrezi, S.edwilsoni, S.ehakernae, Triraphisbillfreelandi, T.billmclarneyi, T.billripplei, T.bobandersoni, T.bobrobbinsi, T.bradzlotnicki, T.brianbrowni, T.brianlaueri, T.briannestjacquesae, T.camilocamargoi, T.carlosherrerai, T.carolinepalmerae, T.charlesmorrisi, T.chigiybinellae, T.christerhanssoni, T.christhompsoni, T.conniebarlowae, T.craigsimonsi, T.defectus Valerio, 2015, T.danielhubi, T.davidduthiei, T.davidwahli, T.federicomatarritai, T.ferrisjabri, T.mariobozai, T.martindohrni, T.matssegnestami, T.mehrdadhajibabaei, T.ollieflinti, T.tildalauerae, Yeliconesdirksteinkei, Y.markmetzi, Y.monserrathvargasae, Y.tricolor Quicke, 1996. Y.woldai Quicke, 1996. The following new combinations are proposed: Neothlipsissmithi (Ashmead), new combination for Microdussmithi Ashmead, 1894; Neothlipsispygmaeus (Enderlein), new combination for Microduspygmaeus Enderlein, 1920; Neothlipsisunicinctus (Ashmead), new combination for Microdusunicinctus Ashmead, 1894; Therophilusanomalus (Bortoni and Penteado-Dias) new combination for Plesiocoelusanomalus Bortoni and Penteado-Dias, 2015; Aerophilusareolatus (Bortoni and Penteado-Dias) new combination for Plesiocoelusareolatus Bortoni and Penteado-Dias, 2015; Pneumagathiserythrogastra (Cameron) new combination for Agathiserythrogastra Cameron, 1905. Dolichozelecitreitarsis (Enderlein), new combination for Paniscozelecitreitarsis Enderlein, 1920. Dolichozelefuscivertex (Enderlein) new combination for Paniscozelefuscivertex Enderlein, 1920. Finally, Bassusbrooksi Sharkey, 1998 is synonymized with Agathiserythrogastra Cameron, 1905; Paniscozelegriseipes Enderlein, 1920 issynonymized with Dolichozelekoebelei Viereck, 1911; Paniscozelecarinifrons Enderlein, 1920 is synonymized with Dolichozelefuscivertex (Enderlein, 1920); and Paniscozelenigricauda Enderlein,1920 is synonymized with Dolichozelequaestor (Fabricius, 1804). (originally described as Ophionquaestor Fabricius, 1804).

Revalidation of Habrobracon brevicornis stat. rest. (Hymenoptera: Braconidae) Based on the CO1, 16S, and 28S Gene Fragments.

Habrobracon hebetor (Say 1836) is an important biological control agent around the world. Many hundreds of papers have been published about its biology. Due to the global distribution of H. hebetor many species have been described over the years in various countries as a sister species to H. hebetor but were eventually synonymized with H. hebetor. One of which is Habrobracon brevicornis (Wesmael 1838). Here, we revalidate the status of H. brevicornis stat. rest. based on molecular data. It remains difficult to discriminate the two species based on morphology data alone, but with molecular data the differentiation is straight forward. An integrative key is provided to distinguish H. hebetor from H. brevicornis.

First record of the genus Phanerotomella (Hymenoptera: Braconidae) from Baltic amber with the description of a new species.

A new but exinct chelonine wasp species, Phanerotomella brevivena Kittel sp. nov. (Braconidae: Cheloninae) of the modern genus Phanerotomella is described from Baltic amber. This new species differs from all other extant Phanerotomella species by having the fore wing vein SR-1 only partially sclerotised. This is the first fossil record of the genus in amber.

A taxonomic review of the parasitoid wasp genera Furcidentia Zettel and Pseudophanerotoma Zettel (Hymenoptera: Braconidae: Cheloninae) from the Neotropics with the description of four new species.

Furcidentia Zettel, 1990 (Braconidae: Cheloninae) from the New World has been recently recognised as a genus separate from Pseudophanerotoma Zettel, 1990. Previously known from three species, three additional new species of Furcidentia are described here from the Neotropical region: F. jenningsi sp. nov. (French Guiana), F. llama sp. nov. (Guatemala), and F. tikalensis sp. nov. (Guatemala). The species F. clypeata Zettel (Ecuador) comb. nov., F. pulchra Zettel (Costa Rica) comb. nov., and F. sharkeyi Zettel (Ecuador) comb. nov. are transferred to Furcidentia. Additionally, one new species of Pseudophanerotoma is described: P. austini sp. nov. (Guatemala), while P. paranaensis (Costa Lima) (Brazil) and P. thapsina (Walley) (Texas, USA) are redescribed and are here recorded from French Guiana for the first time, thus considerably expanding their known distribution. Redescriptions include the description of the male of P. thapsina (Walley) for the first time and of the female of P. paranaensis (Costa Lima). Identification keys to the Neotropical genera of Phanerotomini and the species of Furcidentia and Pseudophanerotoma are provided.

An enigmatic new genus of Hormiinae (Hymenoptera: Braconidae) from South India.

A new Hormiinae genus Indohormius gen. nov. with type species I. keralensis sp. nov. is described and illustrated. Comparison of this genus with some Hormiinae and Rhyssalinae genera are provided. The composition of the subfamily Hormiinae and the position of the new genus on a molecular phylogenetic tree are discussed.

Molecular and morphological phylogenetics of chelonine parasitoid wasps (Hymenoptera: Braconidae), with a critical assessment of divergence time estimations.

Parasitoid wasps of the subfamily Cheloninae are both species rich and poorly known. Although the taxonomy of Cheloninae appears to be relatively stable, there is no clear understanding of relationships among higher-level taxa. We here applied molecular phylogenetic analyses using three markers (COI, EF1α, 28S) and 37 morphological characters to elucidate the evolution and systematics of these wasps. Analyses were based on 83 specimens representing 13 genera. All genera except Ascogaster, Phanerotoma, and Pseudophanerotoma formed monophyletic groups; Furcidentia (stat. rev.) is raised to generic rank. Neither Chelonus (Chelonus) nor Chelonus (Microchelonus) were recovered as monophyletic, but together formed a monophyletic lineage. The tribes Chelonini and Odontosphaeropygini formed monophyletic groups, but the Phanerotomini sensu Zettel and Pseudophanerotomini were retrieved as either para- or polyphyletic. The genera comprising the former subfamily Adeliinae were confirmed as being nested within the Cheloninae. To estimate the age of the subfamily, we used 16 fossil taxa. Three approaches were compared: fixed-rate dating, node dating, and total-evidence dating, with age estimates differing greatly between the three methods. Shortcomings of each approach in relation to our dataset are discussed, and none of the age estimates is deemed sufficiently reliable. Given that most dating studies use a single method only, in most cases without presenting analyses on the sensitivity to priors, it is likely that numerous age estimates in the literature suffer from a similar lack of robustness. We argue for a more rigorous approach to dating analyses and for a faithful presentation of uncertainties in divergence time estimates. Given the results of the phylogenetic analysis the following taxonomic changes are proposed: Furcidentia Zettel (stat. rev.), previously treated as a subgenus of Pseudophanerotoma Zettel is raised to generic rank; Microchelonus Szépligeti (syn. nov.), variously treated by previous authors, is proposed as a junior synonym of Chelonus Jurine; the following subgenera of Microchelonus - Baculonus Braet & van Achterberg (syn. nov.), Carinichelonus Tobias (syn. nov.) and Scabrichelonus He, Chen & van Achterberg (syn. nov.), are proposed as junior synonyms of Chelonus; a number of new species names are proposed due to homonyms resulting from the above changes and these are listed in the paper.

Revision of Megascogaster (Hymenoptera, Braconidae, Cheloninae), with a new species from Sulawesi, Indonesia.

Megascogaster wallacei sp. n. is described from Sulawesi, Indonesia. This is the first representative south of the Philippines and is only the second species of this rare genus described since the type species, M. elongata Baker, 1926. The presence of Megascogaster in Sulawesi, a part of Wallacea, points to a broader distribution of the genus in Southeast Asia. A redescription of M. elongata and a description of the new species M. wallacei are presented along with a key to separate them. 

Remarkable range extension of the previously monotypic braconid genus Wushenia Zettel (Hymenoptera: Braconidae: Cheloninae), with description of a second species from Australia.

The apparently rare chelonine wasp genus Wushenia Zettel was previously known only from a single species Wushenia nana Zettel, collected by Townes at 1150 m from Wushe, Taiwan in 1983. Here we describe a second species, Wushenia australiensis sp. nov. from coastal New South Wales, Australia. This second species extends the known distribution of the genus from the Oriental into the Australasian region, indicating either an extreme disjunct distribution or that Wushenia may also occur on the landmasses inbetween, e.g. the Philippines, Malaysia, Indonesia and/or Papua New Guinea. In addition to a detailed description of the new species, a re-diagnosis of the genus and type species, and a key to species are presented.